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Progress of Plant Breeding in the United States. 



HERBERT J. WEBBER and ERNST A. BESSEY, 

Dioisiou of Vegetable Pliysiology and Pathology. 



Reprint from Yearbook op Department op Agriculture for 1899. 



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7536 



CONTENTS. 



Page. 

Introduction 465 

Early agricultural and horticultural conditions : 466 

Early methods of plant breeding 468 

Evolution of methods of plant breeding during the nineteenth century 470 

Improvements effected during the nineteenth century 475 

Improvement in grapes 475 

Improvement in pears 477 

Improvement in apples 478 

Improvement in plums 479 

Improvement in raspberries 480 

Improvement in blackberries 481 

Improvement in strawberries 482 

Improvement in gooseberries 483 

Improvement in vegetables 483 

Improvement in cereals 486 

Improvement in flowers and ornamental plants 488 

Improvement in nuts 489 

Improvement in cotton 490 



ILLUSTRATIONS. 



PLATES. 

Plate XXXVI. Two prominent early plant breeders 468 

XXX VII. Improvement of the native gooseberry 482 

XXXVIII. Fig. 1. — Improvement of corn by selection. Fig. 2. — Varia- 
tion in seedling pecans 486 

TEXT FIGURES. 

Fig. 22. Plums showing difference between hybrid and parent 479 

23. Hovey's Seedling strawberry 482 

in 



PROGRESS OF PLANT BREEDING IN THE UNITED 

STATES. 

By Herbert J. Webber and Ernst A. Bessey, 
Division of Vegetable Physiology and Pathology. 

INTRODUCTION. 

At the beginning of the nineteenth century a few of the most 
advanced scientific horticulturists were commencing to recognize that 
plants, like animals, are capable of being improved by breeding. 
During the century the knowledge of the factors involved in plant 
breeding gradually increased and became disseminated among prac- 
tical American growers. As a natural consequence of this there 
came to be a better understanding of the methods of plant breeding 
and a greater appreciation of the necessity of securing varieties 
adapted to local conditions, and therefore improved sorts of American 
origin have been gradually but surely supplanting foreign varieties. 

While at the beginning of this period almost all of our cultivated 
fruits, cereals, vegetables, and flowers were of foreign origin, an inspec- 
tion of the present trade lists shows a marked increase of native sorts 
and a corresponding decrease of foreign sorts. In the case of flowers, 
for the production of which artificial conditions largely are main- 
tained, home-produced sorts vie in numbers with those from abroad. 
In cereals and vegetables a majority of the most extensiveby grown 
sorts are of American origin, and in fruits, upon which probably the 
most attention and skill have been brought to bear and the greatest 
stimulus given by well-organized societies, the native sorts have 
almost entirely taken the place of the foreign ones. "In the begin- 
ning of the colonization of this country," writes Bailey, "all the 
varieties of apples were of European origin. But in 1817, over 60 per 
cent of the apples recommended for cultivation here were of American 
origin, that is, American-grown seedlings from the original stock. 
At the present time [1895], fully 90 per cent of the popular apples of 
the Atlantic States are American productions." 

The same increase of American sorts has taken place in the case of 
pears. As early as 1853 Hovey wrote: "It is certainly somewhat 
remarkable, as it is surprising, that, in the course of twenty-four 
years, a larger number of really fine pears have been brought to 
notice, of American origin, than have been introduced from Europe in 
the same time, or we think we might safely add, in the last fifty years." 
While in plums the American seedlings of the European and Japanese 

465 



466 YEARBOOK OF THE DEPARTMENT OF AGRICULTURE. 

species rival the sorts of foreign production, the improved sorts of 
our native species and hybrids of these with the Japan and apricot 
plums are rapidly increasing and will probably soon predominate in 
this industry. In the cultivation of grapes, raspberries, blackberries, 
etc., little advance was made until our native species were taken up 
and improved. All of these have been profoundly modified and 
improved as the result of merely half a century of cultivation and 
breeding. 

It is interesting to note that the present century has witnessed the 
first introduction and wonderful amelioration of some of our now 
most important plants. A striking instance of such a plant is the 
tomato, which is said to have been first brought from Santo Domingo 
to Philadelphia in 1798, but was not sold in the markets of that city, 
according to Manning, until 1829, and did not come into general use 
in the North until some years later. Tomatoes were introduced into 
Salem, Mass. , by an Italian painter, Michelo Corne, in 1802 ; but he was 
said to have had difficulty in persuading the people to eat them. 
They were, however, used as an article of food in New Orleans in 
1812. The wonderful amelioration of the tomato has thus taken 
place wholly within the memory of men now living, and it is not an 
uncommon thing to find aged people, particularly among the pioneers 
of the West, who remember when the tomato was cultivated as an 
ornamental plant, but not thought to be valuable for food. The 
tomato is therefore an excellent illustration of what a century of 
plant breeding may accomplish. 

The flowers now so extensively grown were hardly known a century 
ago, when different varieties were ,just beginning to appear. The mod- 
est chrysanthemum or the carnation of that day would hardly create 
a sensation in our modern flower markets. The immense cut-flower 
trade and the hosts of elegant varieties adapted thereto are the 
results of less than a century of plant breeding. The greenhouse 
has exerted a marked influence on the plants which are thus grown, 
as special varieties are demanded, and the skillful cultivator breeds 
and selects till he secures what is desired. In ail forcing-house 
industries special varieties adapted to this sort of culture have 
sprung up. The changes which have already been wrought are the 
wonder of naturalists and laymen alike, but the end has not yet been 
reached. Everything indicates increased activity in the near future. 
Recent developments, obtained by a few independent experimenters, 
have forcibly called attention to the great improvements which skill 
and patience may achieve in this field, and a renewed interest in such 
matters is very evident throughout the country. 

EARLY AGRICULTURAL AND HORTICULTURAL CONDITIONS. 

In the earty settlement of America agriculture was limited mainly 
to the cultivation of such plants as were known to the settlers in their 



PROGRESS OF PLANT BREEDING. 467 

Old World homes. Each expedition brought seeds and plants to use 
in starting agricultural industries, and subsequent importations of 
desirable varieties continued to be made; hence, the attention of the 
settlers was largely given to testing these experimentally to determine 
their usefulness. The different conditions obtaining in America from 
those found in Europe, from which latter place most of the introduced 
sorts came, rendered the outcome of the early attempts very uncer- 
tain. No exact record of the agricultural development during this 
period exists, but it is probable that the early introduced varieties of 
the various annual crops (cereals, vegetables, etc.) went through a 
gradual evolution and adaptation to conditions by seed selection from 
those plants and strains found to do the best. This selection, which 
the settlers almost certainly exercised, probably did not have any 
definite improvement or change in view other than to secure the best 
and most vigorous seeds. Some of our now most important agricul- 
tural crops, like corn and tobacco, are native American plants, and 
their main improvement consequently dates from the discovery of 
America. In some places, however, the Indians had developed a 
comparatively high state of agriculture, and many sorts of such 
native cultivated plants were obtained from them, as, for example, 
the Golden Sioux, King Philip, and Tuscarora races of field corn. 
An early sweet corn is also recorded as having been obtained from the 
Indians. According to one account it was found and introduced into 
Connecticut by an officer in General Sullivan's expedition against the 
Indians in the Genesee country in 1779. According to another account 
it was introduced into Massachusetts by Capt. Richard Bagnol, of 
Plymouth, who obtained it from the country of the Susquehanna on 
his return from the Sullivan expedition. The Six Nations, against 
which the Sullivan expedition was sent, had made considerable prog- 
ress in agriculture, and are known to have cultivated large fields of 
corn. Besides-this, they are said to have had "gardens of beans, peas, 
turnips, cabbages, melons, carrots, parsnips, and potatoes." 

The earliest attempts at fruit growing in America were mostly 
failures. The varieties grown in early days were nearly all of Euro- 
pean origin.- The recorded history of American horticulture may be 
said not to have begun until the publication of Bernard M'Mahon's 
American Gardener's Calendar in 1806. At this comparatively late 
date native varieties had already become prominent, about 66 per 
cent of the 59 varieties of apples catalogued being of American origin. 
Even at this time, however, very great efforts were still being made 
to extend the range of cultivated products by introductions, the only 
very definite method by which the securing of new sorts was 
attempted. It was a costly experiment, however, and to a great 
extent disappointing. William Kenrick, in a letter to General Dear- 
born (quoted from Robert Manning), says: "From among 150 varie- 
ties imported into Boston by Eben Preble, about 1805, the only 



468 YEARBOOK OF THE DEPARTMENT OF AGRICULTURE. 

additions to the list of desirable kinds were two cherries — the Black 
Tartarian and White Tartarian, and a single pear." 

If we had to-day only the apples and pears known at the beginning 
of the century, the present extensive apple culture of the prairie States 
and the Northwest and the pear culture of the South would be wholly 
impossible. American varieties, the result both of chance discoveries 
and of the most careful and complex methods of plant breeding, have 
almost entirely supplanted the introduced varieties, and are destined 
to become even more important. To-day we look upon plant intro- 
duction as being to a large extent a means to an end. Russian apples 
are being extensively introduced, not wholly with the idea that they 
may become important commercial sorts themselves, but that select 
seedlings from them and hybrids between them and native varieties 
may be obtained, and through these the desired hardy, cold-resistant 
sorts of good quality. 

In the culture of strawberries, raspberries, gooseberries, grapes, etc., 
no material progress was made until the improvement of the native 
species was begun. All these fruits went through an initial stage of 
depending upon foreign varieties, and following this an era of im- 
provement, during which, by careful breeding of the native species 
and infusion into them of the improved European blood by hybridiza- 
tion, strains better adapted to American conditions were obtained. 
This change from an almost total reliance upon introduced varieties 
to a marked supremacy of sorts originated here has taken place almost 
wholly within the past century. 

EARLY METHODS OF PLANT BREEDING. 

The early settlers probably practiced crude selection in growing 
their plants, as anyone, whatever his degree of intelligence, will un- 
consciously do. It is this unconscious selection of individual plants 
through centuries that has led to the important changes which have 
taken place in our principal cultivated plants. So marked has been 
this amelioration that in very many instances the wild forms can not 
now be recognized and are unknown, the most careful study of mod- 
ern scientists having failed to reveal the original types. 

The necessity of having varieties adapted to existing conditions 
was early understood, as was also the necessity of selecting the best 
seed. Manning, in the History of the Massachusetts Horticultural 
Society, says: "And in 1621 the governor requested Massasoit to 
exchange some of their corn, for seed, with ours, that we might judge 
which best agreed with the soil where we lived. The natives were 
acquainted with the advantage of selecting the finest ears of corn for 
seed, and taught the settlers to do the same. They possessed varieties 
adapted to the warmer or colder parts of the country." 

We are inclined to think of plant breeding as based on old and 
well-established laws. The fact is, however, that the fundamental 



Yearbook U. S. Dept of Agriculture, 1899. 



Plate XXXVI. 







mL> 




Two Prominent Early Plant Breeders. 



PROGRESS OF PLANT BREEDING. 469 

principles of intelligent plant breeding were not made known until 
the latter part of the eighteenth century. The sexuality of plants 
was not established until proved experimentally by Camerarius in 
1691, and the first hybrid of which we have any record was not made 
until 1719, when Thomas Fairchild, an English gardener, crossed the 
carnation with the sweet william. 

Our first exact knowledge of hybrids dates from about 1761, when 
Koelreuter began publishing the results of his observations. His 
work was entirely scientific, however, and had but little bearing on 
practical plant breeding, though it served to pave the way for the 
valuable work, soon to follow, of Thomas Andrew Knight (PI. XXXVI), 
the eminent English plant physiologist. The systematic breeding of 
plants may be said to have begun with the work of Knight and Van 
Mons about the beginning of the nineteenth century. Knight was the 
first to show the practical value of crossing and hybridizing in the 
production of plant varieties. In 1806 Knight said: "New varieties 
of every species of fruit will generally be better obtained by intro- 
ducing the farina of one variety of fruit into the blossoms of another, 
than by propagating any from a single kind." 

Another important idea emphasized by Knight, and now quite gen- 
erally accepted, is that one of the principal factors causing or inducing 
variation in plants is an increase of food supply or a modification 
thereof. 

In one sense Knight may be recognized as the father of plant breed- 
ing, no other experimenter having contributed so much toward the 
development of the present system. Of almost equal importance, 
however, was the work of a contemporaneous Belgian horticulturist, 
Jean Baptiste Van Mons, who emphasized mainly the principle of 
selection. His theories were published in various papers, but mostly 
in his Arbres Fruitiers in 1835. His method of obtaining new varie- 
ties of fruit was to gather seeds from young trees in a state of varia- 
tion, taking the fruits before they were fully ripe and allowing them 
to rot, with the idea that this would tend to "subdue or enfeeble" 
the tree, a factor which he thought to be of primary importance. 
These seedlings were then grown in a seed bed until they were Targe 
enough to enable him to judge of their character. He then selected 
the promising ones and planted them in nursery form a few feet apart, 
where they could be fruited on their own roots. When these selected 
seedlings fruited, seeds were taken from the first fruits of the most 
promising and sown, the same process of selection being exercised in 
this second generation, and so on through several or many genera- 
tions of selection till success was attained. The whole process is 
expressed in his own words as follows : "To sow, to re-sow, to sow 
again, to sow perpetually, in short to do nothing but sow, is the prac- 
tice to be pursued, and which can not be departed from; and in short 
this is the whole secret of the art I have employed." 



470 YEARBOOK OF THE DEPARTMENT OP AGRICULTURE. 

As to the theory of selection, time has justified Van Mons's conclu- 
sions, with some modifications, but some of the ideas he advanced 
have been abandoned. He claimed that the older varieties of good 
fruit generally yield inferior seedling sorts, while new inferior varie- 
ties reproduced uninterruptedly for several generations would cer- 
tainly yield good fruit. In these claims the results of recent years 
have shown him to have been somewhat in error. The general theory 
now advanced and used is to select seed for planting from the best 
fruits of the best tree of the best variety. 

The importance of selection in improving varieties was well recog- 
nizee! before the publication of Van Mons's great work above referred 
to, and while the main credit of establishing the principle of selec- 
tion is due to Van Mons, yet other independent workers accomplished 
nearby as important results. Bailey, in his Survival of the Unlike, 
called attention to the work done by Joseph Cooper, of New Jersey, 
in the closing years of the last century. Cooper's observations, re- 
corded in a letter written in 1799 and published in the first volume of 
the Memoirs of the Philadelphia Society for Promoting Agriculture, 
show that he thoroughly understood the action of selection in produc- 
ing changes in varieties. As an • illustration, he says: "A striking 
instance of plants being naturalized happened by Colonel Matlack 
sending some watermelon seed from Georgia, which, he informed me 
by letter, were of superior quality. Knowing that seed from vegeta- 
bles which had grown in more southern climates, required a longer 
summer than what grew here, I gave them the most favorable situa- 
tion, and used glasses to bring them forward, yet very few ripened to 
perfection; but finding them to be as excellent in quality as described, 
I saved seed from those first ripe ; and by continuing that practice four 
or five years, they became as early watermelons as I ever had." 

It is probable that many other advanced horticulturists of that 
period understood and used selection in an intelligent manner. The 
directions given in 1822 by James Thatcher, in his American Orchard- 
ist, for the selection of seeds in attempting to produce improved 
sorts, would be regarded to-day as better than the recommendations 
given by Van Mons. "The seeds for planting," Thatcher wrote, 
"should always be selected from the most highly cultivated fruit, and 
the fairest and ripest specimen of such variety." Thatcher also 
described Knight's method of forcing seedlings into fruit bj^ grafting 
them, and the use of hybridization in the production of varieties. It 
will thus be seen that even at this early date (1822) the fundamental 
principles of plant breeding had apparently become the common prop- 
erty of American agriculturists and horticulturists. 

EVOLUTION OF METHODS OF PLANT BREEDING DURING THE 
NINETEENTH CENTURY. 

It has been seen that the fundamental laws of plant breeding were 
fairly well understood at the beginning of the century and had come 



PROGRESS OF PLANT BREEDING. 471 

to be expounded in horticultural text-books and papers. It must be 
borne in mind, however, that it requires years for scientific principles 
to become thoroughly understood and widely disseminated, so that 
they form a part of common practice. The early native varieties 
were largely chance seedlings, and there now seem to be very many 
choice fruits — pears, apples, grapes, etc. — which originated in this way. 
However, thousands and even millions of worthless wild seedlings, of 
which we have no record, have grown and perished, and in reality 
only one here and there excels and survives. 

During the first fifty years of the century almost the sole method 
of breeding was to select seeds from the best fruits and raise numer- 
ous seedlings, which, when they fruited, were carefully examined, and 
those selected for further propagation which produced desirable fruits, 
of better quality than the parent sorts. One of the earliest syste- 
matic attempts of this sort known to the writers is thus described in 
the Magazine of Horticulture of 1847: "In the fall of 1817, and in the 
following spring, Governor Edwards planted the seeds of pears, with 
the design of obtaining new and superior varieties of this fruit. In 
doing so, he selected the seeds of the best which could be procured, 
including many sorts, but the number was then very limited compared 
with our day." This being one of the first systematic attempts in 
this country to secure improved sorts, it was largely ridiculed. The 
results obtained, however, were of great value, as from some forty 
trees thus produced several fairly good sorts were secured, among 
them being the Calhoun, Elizabeth, Dallas, Henrietta, and Citron. 

In early days, furthermore, up to the fifties, orchards were to some 
extent made up of ungrafted seedlings. When a particularly good 
fruit was produced its seeds were carefully preserved and planted, 
and some varieties were reproduced in the main true to seed. Immense 
numbers of seedling apples were thus grown, and furnished excellent 
opportunity for selection, but only a few produced superior fruit or 
new varieties. In 1845 Rev. Henry Ward Beecher wrote from Indiana 
to the Magazine of Horticulture: "An immense number of seedling 
trees are found in our State. Since the Indiana Horticultural Society 
began to collect specimens of these, more than 150 varieties have been 
sent up for inspection. * * * Of all the number presented, not 
six have vindicated their claims to a name or place — and not more 
than three will probably be known ten years hence." 

Improvement by selection, in the strictest sense of the term, lias 
been employed mostly with annual plants, such as wheat, corn, cotton, 
etc. , and the methods used have been gradually perfected in different 
industries, until in some, as in the sea-island cotton, all growers make 
annual selections with the utmost care to maintain and perfect the 
strain they grow. Very careful methods of selection have also been 
devised to develop and improve corn, and many of the most produc- 
tive and valuable races are the result of continuous selection through 



472 YEARBOOK OP THE DEPARTMENT OF AGRICULTURE. 

numerous generations. In such selection the greatest care is taken 
to secure impregnation with pollen from vigorous, productive plants. 
To insure this the field in which selections are being made is carefully 
gone over when the first silks and tassels begin to appear, and all stalks 
are cut out which are not vigorous and well formed and which do not 
show indications of being productive. By this practice it is brought 
about that fertilization is effected by pollen from vigorous, productive 
stalks only. The final selections are made in the field when the corn 
ripens, the seed ears being taken from the most productive and vigor- 
ous stalks that are true to the type that the breeder is selecting to 
establish. 

In selecting wheat to improve the strain the early attempts were 
mainly confined to simply taking the largest grains — a practice which 
is now recognized as failing in the primary factor of considering the 
productivity and vigor of the individual plant. Many experimenters 
in this country have worked on the improvement of wheat by selec- 
tion, but in general with rather indifferent success. Recently, how- 
ever, Professor Hays, of the Minnesota Agricultural Experiment Sta- 
tion, has used a very careful method of selecting wheat, grown in 
nursery form, which has given valuable results. 

Hybridization and cross-fertilization in improving plants were very 
little utilized in the first half of the century. Knight had started 
the leaven, however, and in some directions it had shown results. 
The idea gradually became current that there was too much chance 
in raising seedlings of unknown parentage. Still, as late as 1857, we 
find the Rural New Yorker giving the following directions in regard 
to raising new fruits: "Eminent pomologists disagree on this subject. 
Our advice, however, is to plant the best seeds of the finest varieties, 
take good care of the plants, and trust to Providence for the result." 
Considerably before this time, however, the most advanced plant 
breeders had given rather different directions. In 1836 A. J. Down- 
ing, one of America's best-known pomologists, wrote : ' 'Assuming Pro- 
fessor Van Mons to be strictly correct, we would suggest that a great 
saving of time and a considerable improvement in quality and vigor, 
might be gained by calling in cross fertilization to the aid of the culti- 
vator, as soon as the fruit of the trees (say the second generation) 
begins to show symptoms of amelioration. By impregnating them 
with the pollen of the finest varieties, we conceive that the next gen- 
eration would produce excellent fruit, and at a saving of twenty or 
thirty years." In 1844 C. M. Hovey, one of the most successful of 
all American horticulturists in the production of improved sorts, 
said with regard to the grape: "Without stopping to institute an 
inquiry into the merits of his [Van Mons] theory, compared with that 
of artificial impregnation, as practiced by Mr. Knight and others, we 
shall recommend to those who would raise seedlings, the importance 
of commencing with the Isabella or Catawba, for one of the parents, 



PROGRESS OF PLANT BREEDING. 473 

and impregnating them with the Sweet Water, Chasselas, or some other 
early foreign variety. The results will be obtained in a shorter 
period, and, we believe, equally as favorable as by the method of suc- 
cessive generations alone." In 1860 Marshall P. "Wilder, in his presi- 
dential address before the American Pomological Society, gave advice 
regarding the origination of varieties in almost exactly the same 
words that might be used to-day: "It was my first, so it shall be my 
continual and last advice; — Plant the most mature and perfect seed of 
the most hardy, vigorous, and valuable varieties, and, as a shorter 
process, ensuring more certain and happy results, cross or hybridize 
your best fruits." 

The first record which we have been able to find of the production 
of a hybrid variety in America is given by Manning, in the History of 
the Massachusetts Horticultural Society, as follows : ' ' Probably the 
first attempt in this country to produce a new fruit by cross-fertiliza- 
tion was by William Prince, who raised the Prince's St. Germain [pear] 
from seed of the old St. Germain impregnated by the White Doyenne, 
about 1806." One of the most successful early attempts in using 
hybridization was by C. M. Hovey, in the improvement of the straw- 
berry, his first hybrid seedlings having been brought to notice in 
1838. He was eminently successful in obtaining good varieties by 
this method, and his success led to the extensive use of hybridization 
in the improvement of this fruit. 

In this connection, it is interesting to note that a striking success 
achieved by any intelligent cultivator in producing valuable varieties 
of any plant has often led to the general adoption of his particular 
methods by other breeders of the same plant. Van Mons's success in 
originating pears by selection led to this method being mainly used in 
breeding this fruit. Allen's success, in 1854, in producing a good 
hybrid grape doubtless stimulated the adoption of this method in 
preference to other methods in improving the grape. 

Since the middle of the century the advance in methods of improv- 
ing plants has been altogether in minor factors. The early hybridizers 
often used a mixture of pollen, believing that it was possible for the 
same seed to be influenced by pollen from several varieties, or species. 
The details of the process of fecundation were not well worked out at 
that time, and it is not surprising that early experimenters frequently 
erred in their conclusions and were thus led to pursue false methods. 
With the gradual increase in knowledge of the methods of fecundation 
the idea of the effectiveness of using two kinds of pollen at the same 
time was abandoned, and in casting about for other methods of secur- 
ing the results sought growers evidently began the practical use of 
compound hybrids, as the method came into practice about this time. 
Numerous hybrid rhododendrons, begonias, etc., contain the blood of 
several species, mingled with the definite idea of securing in the off- 
spring certain characteristics from each parent. Compound hybrids 



474 YEARBOOK OF THE DEPARTMENT OF AGRICULTURE. 

have been particularly valuable in grapes, among the numerous excel- 
lent sorts of such hybrids being Lady Washington, Brighton, and 
Brilliant. 

Another important factor in the application of hybridization to 
securing improved strains, and one which has but very recently become 
prominent, is the securing of what have been termed dilute hybrids, 
that is, hybrids containing more blood of one variety than of the other. 
If in any hybrid the character of one of the parents is found to be too 
pronounced to give a successful combination, it is crossed with the 
other xDarent, the result being a three-fourths hybrid, that is, a hybrid 
deriving three-fourths of its characters from one of the original parents 
and one-fourth from the other. 

The value of selecting distinct parents and introducing new species 
into combination with old ones was early recognized, but mainly 
among florists, where a change of color was desired. In 1836 Hovey 
called attention to the change in color produced in calceolarias by the 
introduction of a different-colored species. "It was not until the 
introduction of a purple species, C. purpurea, in 1827," writes Hovey, 
"that any variation took place in the color of the flowers; the pre- 
viously introduced ones being yellow, of course no other shade was 
produced until the impregnation of the former with the latter. At 
the present time, however, plants are to be found of almost every tint, 
from the palest yellow to deep orange, and from light red to bright 
scarlet, as also, two or three of these shades distinct in the same 
flower." The results of more recent work have emphasized the 
importance of using very distinct parents when marked changes or 
new creations are desired. The improved strains of begonias and 
roses resulting from the introduction of Begonia socotrana, Rosa 
rugosa, and R. wichuraiana, and Burbank's walnut hybrids (crosses 
of Juglans californica, J. regia, and J. nigra) illustrate the impor- 
tance of this practice. This has led in recent years to the extensive 
introduction of and experiments with various wild species of common 
cultivated plants, and the field here opened to the horticulturists and 
florists is one of promise. 

The importance of growing hybrids through several or at least two 
generations, in order to secure greater variation, particularly where 
the hybrid is from widely distinct parents, was scientifically demon- 
strated by Naudin and Nageli in 1865. The practical importance of 
this discovery, however, has come to be thoroughly understood and 
appreciated by American plant breeders only in the closing years of 
the century. 

In very recent years there has been much discussion of the ques- 
tion of the improvement of certain cultivated plants by selection of 
the vegetative parts used in propagation. It seems to have been 
proved beyond question that certain plants can be greatly modified in 
this way, particularly as to vigor and productiveness. This method 



PROGRESS OF PLANT BREEDING. 475 

of improvement seems likely to play a very important part in the 
future by aiding to secure strains of standard sorts suitable for 
growth in special localities and varying but slightly from the original 
varieties. 

IMPROVEMENTS EFFECTED DURING THE NINETEENTH CENTURY. 

In the present paper it is possible to call attention to only a few of 
the most important improvements illustrative of the advances made 
in certain fields of agriculture and horticulture. In early days, as 
previously indicated, the majority of the native varieties introduced 
were merely chance seedlings, which grew uncared for until their 
good qualities were discovered, when they were brought into cultiva- 
tion. The sorts obtained in this way are not primarily due to plant 
breeding, being simply the result of intelligent choice of chance-sown 
plants, yet some of these varieties have had a marked influence on 
the development of certain industries. Of far greater importance, 
however, has been the introduction of varieties which have been pro- 
duced by careful methods of selection, carried through from one to 
many generations. 

Hybridization also has already had a very marked effect in the 
development of many cultivated plants, and in the future it will 
doubtless be extensively utilized in securing desired modifications. 

IMPROVEMENT IN GRAPES. 

The grape has been very much improved by American cultivators 
and furnishes an excellent illustration of the great amelioration which 
may be obtained in a comparatively short period. For many years 
after the settlement of America the only grapes cultivated were of 
European origin. Numerous trials, however, proved that these were 
not hardy in Eastern America, and that they soon succumbed to 
attacks of Phylloxera and other diseases. Curiously enough the 
native American grapes, which were found in great abundance 
throughout the eastern part of the country and attracted consider- 
able attention, were for years neglected, and it was only after the 
failure of the European sorts had been demonstrated that the native 
sorts were brought into cultivation. The first of these to attain 
prominence was the now famous Catawba, which was found wild in 
North Carolina in 1802, and was brought into general notice by Maj. 
John Adluni, of Georgetown, D. C. A few j^ears later the Isabella, 
another wild grape, was introduced, and after the success of these 
two sorts had been demonstrated many other wild forms were brought 
into cultivation. 

Apparently very little systematic effort was made to improve the 
grape until the appearance of Pond's Seedling in 1835. The time of 
its introduction is worthy of notice as being the beginning of a period 
of planting seeds of the native species for the purpose of making 
selections. 



476 YEARBOOK OF THE DEPARTMENT OF AGRICULTURE. 

The greatest advance in grape culture in this country is without 
doubt due to the famous Concord, which was also produced by selec- 
tion. About the year 1840, Mr. Ephraim Bull, of Concord, Mass. 
(PI. XXXVI), found growing on his grounds a wild grapevine, which 
was apparently a seedling from some wild grapes that had been 
scattered about his place by boys the preceding year. He took up the 
vine and moved it to his garden, giving it good care until it fruited 
in 1843, the fruit, which was of good quality, ripening as early as the 
latter part of August. He was so impressed with the superior quality 
of this fruit and the lack of foxy flavor that ' ' the idea at once occurred 
to him that another generation would be a still greater improvement. " 
Following this out, he planted seeds of this grape, obtaining a num- 
ber of seedlings. One of these, which fruited first in 1849, was so 
markedly superior to the others that it alone was preserved, later being 
named the Concord. This grape, because of its vigor, productiveness, 
and fine quality, at once became very popular. Not only has the variety 
proved of great value itself, but it has been the parent of a great num- 
ber of varieties, many of them of considerable merit. Probably the 
best known of these are the Worden and Moore's Early Mr. Bull con- 
tinued to plant seeds of the Concord year after year until he had pro- 
duced over twenty-two thousand seedlings, but of these there were 
only twentj 7 -one which he recommended for cultivation, and none of 
them have become as popular as the parent variety. 

In the last twenty years very many varieties of the grape have 
been produced, but no select seedlings of striking importance have 
appeared, the good new varieties being mainly hybrids. In this con- 
nection it is worthy of notice that the grape owes more to hybridiza- 
tion than does any other fruit. The Delaware grape, which is even 
yet a standard of excellence, is probably a natural hybrid, containing 
some blood of the fine European grape. This was found in a garden 
of foreign grapes in New Jersey about the year 1850, but received its 
name from Delaware, the Ohio town in which it was first brought to 
general notice. It is undoubtedly the best of our chance seedlings, 
and was the last introduced that proved of much merit. 

Many of our most widely cultivated varieties of grapes, such as the 
Salem, Niagara, Brilliant, etc., which are common sorts in the mar- 
kets, are the results of careful hybridization. The first hybrid grape 
produced in this country, known as Allen's Hybrid, was introduced in 
1854, and was a cross of the Isabella with a European variety, sup- 
posedly Golden Chasselas. This is the epoch-making grape as far as 
hybrids are concerned. It was regarded with much interest because 
of its fine quality and appearance, and while the bright hopes regard- 
ing it were never realized, it was of the greatest importance, as it 
served to stimulate the improvement of grapes by hybridization. 
Shortly after this E. S. Rogers, of Roxbury, Mass., began introducing 
his new hybrid varieties, the first being sent out in 1856. His Salem 



PROGRESS OF PLANT BREEDING. 477 

is an excellent chestnut-colored sort, and is probably the most exten- 
sively grown of any hybrid grape. Rogers was closely followed by 
Ricketts, Burr, Caywood, Moore, Rommel, Stayman, and several 
others, who were very active in the production of new sorts, mainly 
hybrids, and more recently still, by T. V. Munson, of Denison, Tex., 
who has probably conducted the work on a more extensive scale than 
any other experimenter in this field. Munson has already sent out 
thirty-six new varieties, for the most part hybrids, and is still actively 
engaged in the work. 

A tabulation of the grapes described in Bush & Son & Meissner's 
Grape Grower's Manual shows that, of 554 varieties described, 287 are 
hybrids, 141 select seedlings, 57 chance seedlings, 68 of unknown ori- 
gin, and 1 a sport. Considering those of known parentage, 59 per 
cent are hybrids, 29 per cent select seedlings, and 12 per cent chance 
seedlings. These figures show the marvelous extent to which hybrid- 
ization has affected the improvement of the grape. 

IMPROVEMENT IN PEARS. 

One of the first native varieties of pears to be introduced was the 
Seckel, which has remained to the present time our standard of excel- 
lence. It was found near Philadelphia during the eighteenth cen- 
tury, apparently being a chance seedling. Many other early native 
varieties introduced were obtained in this way, among them Tj'son, 
Andrews, and the Columbia Virgoulouse, the last named remaining 
a xjopular pear for a considerable time. It was not long, however, 
before the practice of planting seeds of the best fruits and selecting 
from the resulting seedlings came to be adopted in the improvement 
of the pear. One of the first attempts of this kind to attract attention 
was that of Governor Edwards, of Connecticut, as mentioned elsewhere. 
Probably the most systematic and successful attempt at growing seed- 
lings for selection was that made by Mr. Dana, of Massachusetts. He 
planted seeds of the best varieties and raised five or six thousand 
seedlings, from which he obtained many good varieties, the best being- 
Dana's Hovey, introduced about 1860. It is worthy of note that Dana 
always planted the seeds of the best varieties, a practice directly oppo- 
site to Van Mons*s theory, and yet succeeded in producing many good 
sorts. 

The pear owes but little of its development to artificially produced 
hybrids, and yet in no other fruit have hybrids played such an impor- 
tant role. The Kieffer, Le Conte, and Garber, all widely-grown com- 
mercial pears, through which this industry has been greatly extended, 
are naturally-produced hybrids of the European pear and the Chinese 
sand pear. 

The European pear, noted for its excellent qualitj 7 , succeeds admir- 
ably on the Pacific coast, but has never proved wholly satisfactory in 
the Eastern States, and can not be successfully grown on a commercial 

2078 2 



478 YEARBOOK OF THE DEPARTMENT OF AGRICULTURE. 

scale south of Virginia. The Chinese sand i^ear comes from a region 
having climatic conditions very similar to those of the Eastern and 
Southern States, and thus finds here a congenial home. The fruit 
is of poor quality, however, and the variety is grown only as an orna- 
mental tree and for stocks on which to hud other sorts. The Kieffer 
and Le Conte are both seedlings of the Chinese sand pear, and from 
their characters show that the seeds from which they grew must have 
been accidentally crossed with the pollen of some good variety of the 
European pear. It is probably to the father parent, the European 
pear, that is due the improved quality of the fruit, while the vigor and 
adaptability to growth in warm climates evidently come from the 
mother parent, the sand pear; These hybrid sorts practically revolu- 
tionized pear culture in the Eastern United States, extending the limit 
of profitable commercial pear growing several hundred miles south- 
ward. From Virginia to Florida these varieties grow luxuriantly and 
have practically driven out all other sorts. Even as far north as 
Philadelphia the Kieffer is by far the most important commercial 
variety. 

IMPROVEMENT IN APPLES. 

Among apples, as in the case of pears, the variety that is consid- 
ered to be a standard of excellence, the famous Newtown Pippin, was 
obtained as a chance seedling. It was introduced to notice about 
two hundred years ago. The Baldwin apple, which has exercised 
such an important influence on the apple industry, was also a chance 
seedling, which sprang up about 1742 on the farm of Mr. John Ball, 
in eastern Massachusetts, and was brought into general notice by a 
Colonel Baldwin, from whom it took its name. This apple proved to 
be of such importance that its origin has recently been commemo- 
rated by the erection of a monument on the spot where the original 
tree stood. 

Many other chance seedlings have proved to be of great value, but 
a large proportion of the varieties of most importance, obtained dur- 
ing the nineteenth century, are the results of selection either of 
seedlings grown for the purpose or from seedling orchards. Of these 
may be mentioned the Northern Spy, originated in New York about 
1800; the Jonathan, introduced in 1829; the Summer Bellflower, and 
many others. 

One of the important problems which has recently taxed the skill 
of apple breeders has been to secure varieties suitable for growth in 
the northwest prairie region. The Wealthy apple, the first variety 
to meet this condition, furnishes one of the most striking examples of 
improvement in apples effected by planting numerous seeds and 
selecting from the seedlings. About the year 1855 Mr. Peter M. 
Gideon, of Minnesota, began fruit culture, planting fruit trees of 
various kinds, among them thirty named varieties of apples, and also 
a bushel of apple seeds. Each succeeding year for nine years he 



PROGRESS OF PLANT BREEDING. 



479 



planted more trees and also enough seeds to produce about a thou- 
sand trees each year, hut the cold winters kept killing them off! until 
at the end of the ten years there was left only one small seedling crab. 
All of Mr. Gideon's neighbors gave up the attempt to grow fruit, 
characterizing it as an impossibility, and urged him to do the same, 
but he persisted and sent to Bangor, Me., for scions and seeds. From 
the seeds of the Cherry Crab thus obtained one seedling proved hardy 
and was named the " Wealthy." Upon these varieties the apple cul- 
ture of the northern Mississippi River region has been built. Within 
very recent years there has been great activity in hybridizing our 
different varieties of the apple with the varieties of Russian apples 
recently introduced and with the native wild crab, the object being 
to obtain hardier varieties. This line of experiment, started in the 
closing years of the century, "will probably in a few years yield results 
of the greatest practical value. 

IMPROVEMENT IN PLUMS. 

For many years plum culture in America was almost entirely limited 
to the cultivation of introduced A^arieties of the European plum, but 




Fig. 22.— Plums showing difference between hybrid and parent: Hybrid plum Golden at right 
and mother parent Robinson at left, natural size (after Burbank). 

little attention being given to the origination of native sorts, as, in the 
main, the finer foreign sorts succeed fairly well in the limited area in 
which the European plum can be grown. Nevertheless, some attempts 
were made, and in Canada Henry Corse grew thousands of seedlings 



•480 YEARBOOK OF THE DEPARTMENT OF AGRICULTURE. 

for several years previous to 1840 with the hope of procuring something 
which would excel existing varieties. From this great number he 
selected several which promised well, and to four, of whose excellent 
quality there was no doubt, he gave the names Dictator, Victoria, 
Colonel Wetherell, and Nota Bena. About the middle of the century 
the value of our native species came to be recognized. Many selected 
chance seedlings were brought into cultivation, and it is to these that 
we owe the development of our native species of plums, nearly all the 
best-known varieties of which were obtained in this manner. 

It is only recently that any attempt has been made to improve our 
plum's by artificial hybridization, and this attempt has been brought 
about mainly by the introduction of the Japan plum (Prunus tri- 
flora), which has entered into most of the valuable combinations 
thus far produced. The first Japan plum grown in this country, the 
Kelsey, was not introduced until 1870. The great activity in intro- 
ducing the Japanese varieties and crossing them with American sorts 
did not begin, however, until several years later, but, according to 
Professor Waugh, the Japan plum alreacty constitutes one parent of 
twenty-seven hybrids which have been found valuable and named. 
The introduction of this plum and its use in hybridization bids fair to 
be of the greatest importance to the plum industry . Luther Burbank, 
of California, was the pioneer in plum hybridization, and has pro- 
duced very many valuable sorts, such as the Golden (fig. 22), Juicy, 
and America (crosses of Robinson with Botan). The apricot plum, 
another species, has also been used a number of times by American 
experimenters in crossing with the Japan plum, and has yielded such 
fine combinations as the Climax, Chalco, Late Conical, and probably 
the Wickson, all of which were produced by Burbank. Some valu- 
able hybrids of our native species have also been produced, but they 
are not so promising as hybrids witli the Japan plums. 

IMPROVEMENT IN RASPBERRIES. 

The varieties of raspberries cultivated in this country are almost 
entirely of the native species, it having been found difficult to grow 
the European varieties. Accordingly, we find that our first varieties 
are derived 'mostly from wild plants picked up in the woods and the 
fields and brought into cultivation. Among those thus cultivated, 
probably the first to be named and generally distributed was the 
so-called English Red, which was really a native American variety. 
Among other chance seedlings are the Ohio Everbearing, Catawissa, 
and Cuthbert. The last named was found growing in a garden in 
Riverdale, N. Y., in the latter part of the seventies, and soon became 
a popular sort. 

The systematic improvement of the raspberry by growing seedlings 
for selection was much retarded by the earlier growers of this fruit 
attempting to make use of the European instead of the native species. 



PROGRESS OF PLANT BREEDING. 481 

In the meantime, however, many wild plants of the American species 
were domesticated on farms and in gardens. Among the early experi- 
menters with this fruit was Dr. Brinckle, of Philadelphia, who pro- 
duced a great many varieties, hut only one which proved important. 
This was the Brinckle's Orange, produced in 1844, from an English sort 
known as Dyark's Seedling. It has proved to be a very popular berry, 
and has been widely grown, being one of the very few varieties of the 
European species to prove hardy in America. Soon there appeared 
other varieties, many of them being seedlings of foreign sorts, but prob- 
ably in many cases accidentally crossed with the native species. Most 
of the varieties now grown, however, are improved varieties of the 
American species. 

In the last quarter of the century several valuable hybrids have been 
introduced which have become popular sorts. Among these may be 
mentioned the Dictator (Gregg crossed with Schaffer) and the Caroline 
(Brinckle's Orange crossed with Black Cap). 

IMPROVEMENT IN BLACKBERRIES. 

The blackberry, as a cultivated plant, is entirely an American pro- 
duction, and we owe nothing to the European plant breeders so far as 
it is concerned. All the earlier varieties were merely wild plants 
taken up and set out in the garden. One of the first attempts to 
improve the blackberry was that by Mr. Lovett, of Massachusetts, who 
for many years attempted to find good plants and bring them into cul- 
tivation. It was not until 1850, however, that the Dorchester, the first 
variety to be named, was introduced. In 1854 a berry was introduced 
that marked an epoch in blackberry culture, and showed what the 
fruit was capable of becoming. This was the Lawton, or New Rochelle, 
as it is often called. It was found by the roadside near New Rochelle, 
N. Y., and was introduced by Mr. Lawton. This berry long remained 
popular, but its place was finally taken by Wilson's Early, also found 
as a wild plant. 

The culture of the blackberry is still in its infancy, and compara- 
tively little attention has been given to its improvement. Quite a 
number of hybrid varieties, such as Iceberg, Autumn King, Minne- 
waska, etc., have been introduced, but none have as yet become very 
well known. 

The raspberry and blackberry have been repeatedly hybridized by 
experimenters like Burbank and Carman, and some suggestive results 
obtained. Burbank's series of raspberry-blackberry hybrids are in 
many respects the most remarkable ever produced between distinct 
species. The most noteworthy of these hybrids are Primus (Western 
dewberry crossed with Siberian raspberry), Paradox (Crystal White 
blackberry crossed with Schaffer raspberry), and Humboldt (Improved 
California Wild dewberry crossed with Cuthbert raspberry). Bur- 
bank, in speaking of the Primus, says: "It is also remarkable that 



482 YEARBOOK OF THE DEPARTMENT OF AGRICULTURE. 

the hybrid should ripen its fruit several weeks before either of the 
parents, and excel them much in productiveness and size of fruit, 
though retaining the general appearance and combined flavors of 
both." The Paradox was the only one retained out of some forty 
thousand hybrid seedlings. 



IMPROVEMENT IN STRAWBERRIES. 

Strawberry culture in this country was conducted on a small scale 
at first because no varieties well suited to the climatic conditions 
were known. Many were tried without success, especially the Keen's 
Seedling, which was represented as very promising; but while it was 
an exceedingly valuable berry in England, its place of origin, it failed 
to fulfill the expectations of those who imported it into this country. 
After cultivating this and many other sorts of more or less note, Mr. 
C. M. Hovey, an eminent American poinologist, became satisfied that 
there existed in this country at that time no variety possessing the 

qualities necessary to make its 
cultivation profitable. "There 
seemed to be wanting," says 
Hovey, "a variety combining the 
qualities of two or more of these, 
and we set out upon the experi- 
ment of attaining this desirable 
result, determined, if time would 
allow, to pursue it until our ob- 
ject was accomplished. " He pre- 
pared plants of seven distinct va- 
rieties, and in 1833 made six series 
of crosses, having first carefully 
removed the stamens from the 
flowers to be pollinated, so as to prevent self-fertilization. The fol- 
lowing year the resulting seeds were planted and produced plants of 
very varied appearance and characteristics. Only a few of these sur- 
passed the best of the parent varieties, but one in the size and num- 
ber of its berries, as well as in fine flavor, excelled anything known in 
this country. This plant was kept under observation six years, at the 
end of which period, having fulfilled the expectations of its producer, 
it was put on the market under the name of Hovey's Seedling (fig. 23). 
The effect of the production of this berry was truly wonderful, and 
resulted in making strawberry culture popular and profitable. Other 
experimenters began to make crosses and to grow new sorts, but in 
spite of the almost innumerable varieties thus produced Hovey's Seed- 
ling remained the leading berry for almost thirty years. It is worthy 
of note that although it was probably the most famous variety ever 
produced, it is now extinct, it being impossible to obtain typical plants. 




Fig. 83.— Hovey's Seedling strawberry) half natu 
ral size (from the Magazine of Horticulture). 



Yearbook U. S. Dept of Agriculture, 1899 



Plate XXXVII. 




Improvement of the Native Gooseberry: 1, Ribes oxyacanthoides, Wild Form; 
2, Houghton Gooseberry, Seedling of the Wild Form ; 3, Downing Gooseberry, 
Seedling of the Houghton. (All Natural Size, Adapted from Bailey. i 



PROGRESS OF PLANT BREEDING. 483 

The Wilson, which supplanted it, is itself being gradually supplanted 
by other sorts, and will probably eventually disappear. Among the 
varieties produced from Hovey's Seedling may be mentioned Moya- 
mensing Pine, which in 1849 was awarded the prize offered by the 
Philadelphia Horticultural Society for the best new berry. This was 
in turn the parent of many other varieties, some of which were of 
considerable merit. 

Hybridization has been the favorite method of producing new 
varieties of strawberries, perhaps because the first successful variety 
was obtained in this way. Among the recent hybrids may be men- 
tioned the Hunn, which also illustrates the difficulty of systematic 
breeding, it being the only one deemed worthy of preservation out of 
about seventeen hundred hybrid seedlings tested. 

IMPROVEMENT IN GOOSEBERRIES. 

As in the case of most other fruits, the first varieties of gooseberries 
grown in this country were of foreign origin. However, these mil- 
dewed very badly, especially when their cultivation became more gen- 
eral; hence, in the course of time the growing of this fruit was almost 
totally abandoned. Soon the native species began to attract atten- 
tion, however, and one of the first varieties to be described was 
Houghton's Seedling, produced near Lynn, Mass., about 1845, from 
the wild gooseberry. A few years later Mr. Downing, of New York, 
produced from this already popular variety the Downing, a seedling 
which has since become extremely popular. The Houghton and Down- 
ing, compared with the wild type from which they sprang, furnish an 
interesting illustration of the evolution of a native wild plant. (PI. 
XXXVII.) Since then a number of seedling varieties of good quality 
have been produced, and have come to be quite extensively culti- 
vated. However, now that the use of fungicides has become general, 
the English varieties are again coming into cultivation, and it is 
still a question whether the advantage gained b} r the American varie- 
ties, owing to the exclusion of the former by mildew for so many years, 
will enable the American sorts to retain their supremacy. 

IMPROVEMENT IN VEGETABLES. 

The tomato. — The tomato illustrates well what can be accom- 
plished by careful breeding. In the early part of the century the 
races of tomatoes had mostly small and lobed fruits, but in the course 
of fifty years or more of selection the type has. changed until the fruit 
is now large and smooth and the habit of the plant very different. 
As in the case of the strawberry, the first great advance in the devel- 
opment of the tomato in this country was made by hybridization. 
The Trophy tomato, introduced by the late Colonel Waring, was the 
first of our modern, smooth, round tomatoes, and its production ami 



484 YEARBOOK OF THE DEPARTMENT OF AGRICULTURE. 

advertising, probably more than anything else, served to make the 
tomato a popular garden vegetable. "The Trophy tomato," in the 
words of Colonel Waring, " is a product of crossing and careful cul- 
tivation by Dr. Hand, of Baltimore County, Md., who began his 
work in connection with it about 1850. He crossed the small, smooth 
'Love Apple,' which was filled with juice and seeds, with the com- 
pound, convoluted tomato of that period. This latter was practically 
four or five separate fruits packed together in one, with the skin 
running far into the convolutions. He succeeded in putting the 
solid mass of this compound growth into the smooth skin of the Love 
Apple, and then, by careful selection, year after year, increased its 
size and the solidity of its contents until it became a mass of flesh 
interspersed with small seed cells." The Trophy remained for a 
number of years the principal race on the market, but was finally 
superseded by others bearing larger and better fruits, in the produc- 
tion of which hybridization played an important part. The effect of 
selection in recent years is illustrated in the production of the Para- 
gon by A. W. Livingston. In passing through a field of tomatoes, 
he selected one plant because of the uniformly smooth fruits and 
because of its being very prolific. The seeds from this plant were 
sown the next year, and the stock of seeds for planting was saved 
from the earliest and best specimens. By continuing this process for 
five years the Paragon was produced. The Acme, Perfection, and 
many other races were originated in a similar manner. 

The potato. — The potato has long been the subject of more or less 
systematic improvement in this country. According to Bailey, even 
as early as the end of the last century, Joseph Cooper made ' ' success- 
ful experiments in keeping and improving strains of the potato." 
There is a record in 1835 of the production of anew variety called Per- 
kin's Seedling, originated by planting a seed ball a year or two before. 
In 1841 the Pollard, a seedling of the Chenango, was introduced, but 
the most popular potato originated during this period was the Mercer, 
which was also a select seedling. For a good many years after this it 
was a general practice to plant potato seeds to produce new varieties, 
but these for the most part remained known only locally. 

The introduction of new wild strains from South America marked 
the beginning of a very distinct epoch in the culture of the potato. 
About 1850, or possibly two or three years earlier, a Mr. Goodrich 
began experiments with a view of improving the potato, using the 
varieties known as the Wild Peruvian and the Rough Purple Chili, 
which were either direct importations from South America or but 
slightly improved. He grew seedlings of these varieties for fifteen 
years, obtaining over sixteen thousand, but considered only ten of 
this number worthy of cultivation. The best two of these were the 
Cuzco, a seedling of the Wild Peruvian, and the Garnet Chili, from the 
Rough Purple Chili. Later the Cuzco gave rise to several fairly 



PROGRESS OF PLANT BREEDING. 485 

valuable vai-ieties, but it was surpassed by the Garnet Chili, from 
which several of our best-known varieties have been produced. 

In 1860 or 1861 a grower of the Garnet Chili preserved a seed ball 
of this variety, pinning it up against his window until it was old and 
dry, when, fortunately for the potato industry of the United States, 
he gave it to Mr. Albert Breese, of Vermont. Mr. Breese planted the 
seeds and obtained widely varying plants, some producing many 
tubers and others but few, while there was no uniformity in their size 
or shape, some being large and others small, some round and others 
elongated. Seven of the plants proved to be of exceedingly good 
quality, but one of these, an early sort, far surpassed the others and 
was named the Early Rose. When this potato was put on the market 
a few years later, it commanded almost fabulous prices, and in a few 
years became the leading variety in America, a position which it still 
retains over a considerable part of the country. The other varieties 
of similar origin also became quite popular, and soon the old sorts 
were completely abandoned. In fact, there can be found in the cata- 
logues of varieties grown at the present time scarcely a single variety 
popular forty years ago. 

In the production of many sorts, such as the White Elephant, Snow- 
flake, Nebula, etc. , hybridization has been used, but so far none of 
the varieties thus produced have proved as valuable as the Early 
Rose. 

The potato has been also somewhat improved by the selection of its 
tubers. Thus, when the Early Ohio was introduced, a careful selec- 
tion was made of the "medium-sized, well-ripened tubers of a desired 
shape," with the result, according to C. L. Allen, of "fully a week's 
gain in earliness; a great increase in productiveness, with a marked 
decrease in the quantity of vines." A few of our well-known varie- 
ties originated as bud sports from the tubers ; for example, Thor- 
burn's Late Rose from tubers of the Early Rose. 

The garden pea. — The garden pea furnishes an example of great 
improvement produced largely by hybridization, the most marked 
result obtained in this country being the production of the dwarf pea 
American Wonder about the year 1880, up to which time the varieties 
grown were almost all of foreign origin. This variety was the result 
of a combination of McLean's Little Gem and the Champion of Eng- 
land. The former long stood at the head of the dwarf peas, but was 
unproductive; the latter at the time the cross was made was consid- 
ered the best in quality and the most productive of the tall peas. The 
American Agriculturist says: "We look upon the production of this 
pea as one of the most important steps made of late in its department 
of horticulture." 

The squash. — It is interesting to note that among squashes, which 
hybridize so readily, the Butman, originated by Mr. Clarendon Bat- 
man, of Maine, about 1875, was the result of crossing the Hubbard 



486 YEARBOOK OF THE DEPARTMENT OF AGRICULTURE. 

with a Japanese race and of several years of careful selection. This, 
according to Mr. James J. H. Gregory, was the first instance of a race of 
squashes produced in America, all of our standard races previous to 
the production of this one having been originated abroad. 

IMPROVEMENT IN CEREALS. 

Corn. — Corn has probably been more or less the subject of improve- 
ment by selection ever since it was first cultivated, and it is a general 
practice among farmers who grow their own seed to select the best 
ears for the next year's planting. There have been some experiment- 
ers, however, who have given special attention to its improvement, 
among the earliest being Mr. J. S. Learning, who began in the early 
fifties by going through his fields — then producing an ordinary, not 
very prolific, yellow corn^-and selecting seed from the best-formed 
plants bearing two or three well-formed oars. In this way, by a con- 
tinuous selection extending over thirty j^ears, the famous Learning 
corn was produced and kept up to its standard. About fifteen years 
later Mr. James Riley, of Indiana, also began the careful selection of 
corn, taking a fine white sort as the original. He used essentially the 
method followed by Learning, but in addition went through the fields 
just as the tassels were appearing and cut out all imperfect and bar- 
ren stalks. He selected seed for the next year's planting from the 
finest stalks and the best and most evenly developed ears. By con- 
tinuing this selection for several years he produced the Boone Countj^ 
White (PI. XXXVIII, fig. 1), which has given noteworthy yields at 
the Illinois experiment station. 

Corn has been greatly modified and improved by hybridization, 
but no improvement stands out as marking a distinct epoch. The 
earliest account of a new race being originated by hybridization 
which has come under the notice of the writers is that of the Smith's 
Early White, described in a letter by Dr. Gideon B. Smith, in the 
Albany Cultivator for 1838, the experiments being said to have been 
started some ten or twelve years earlier. It was the result of a cross 
between the Tuscarora and the Sioux. Dr. Smith's discussion shows 
that the results to be expected from crossing different races of corn 
were thoroughly understood even then. The original Old Colonj^ 
sweet corn, a race originated about 1849, and extensively cultivated 
for years, was one of the first and best of the sweet-corn hybrids. 

The ease with which corn hybridizes naturally in the field has led 
to great mixing, and doubtless many forms now cultivated are selected 
types of such accidental crosses. Very many of the best races, how- 
ever, were originated as carefully produced hybrids. 

Wheat. — The early races of wheat grown in this country were, as 
was the case with almost all our cultivated plants, of foreign origin, 
and even now a great many sorts are being imported, especially from 
Russia. A large number, however, have had their origin in America, 



Yeaibook U. S. Dept. of Agriculture, 1899. 



Plate XXXVIII. 




Fig. 1 .—Improvement of Corn by Selection: Boone County White Corn on Left 
and Original Type from which it was Developed by Selection on Right. 




Fig. 2.— Variation in Seedling Pecans: Frotscher Pecan on Left and Two Seed- 
lings from it, Showing Variations in Thickness of Shell, Size, etc. (Natural 
Size.) 



PROGRESS OF PLANT BREEDING. 487 

the first of these being mainly such as originated in fields of wheat or 
from chance-sown seeds, which, owing to their differences from other 
wheat, were preserved and perpetuated. Such, for example, were the 
Tappahannock, found in Virginia in 1854, and the famous Fultz wheat, 
found in a field of Lancaster Red wheat in Pennsylvania in 1862 by 
a Mi\ Abraham Fultz. Mr. Fultz was attracted by some beautiful 
heads of smooth wheat, which he saved and planted by themselves, and 
from these the new race was developed. 

But little attention has been given to the systematic growing of 
wheat for selection until quite recently. The most important experi- 
ments of this kind in the United States are those by Prof. W. M. 
Hays, of the Minnesota experiment station, which are still in prog- 
ress. From the year 1888 up to the present year 552 different races 
have been tested, from which eight were finally selected as worthy of 
preservation. From these eight, selection experiments were started 
in 1892, and as a result, even at the end of the first year, four of the 
best eight new strains surpassed in yield and in some other qualities 
the best four of the old varieties. Though it is too early yet to give 
more definite results, it is evident that the use of selection is very 
promising as regards the improvement of even the best races. 

Within recent years considerable attention has also been given to 
hybridization, and many valuable hybrids have found places in our 
lists of important races. Attention has been directed mainly to in- 
creasing the yield by crossing different strains and to securing earlier 
and hardier sorts. Among the earlier experimenters in this field 
Arnold and Pringle were eminently successful. Arnold's Hybrid No. 
9, a cross of Michigan Amber with White Soules, has in some places 
given good results. Pringle's Defiance, said to be "a hybrid of a 
white wheat common in California upon an Eastern club variety," 
has proved very valuable in California, Colorado, and other places. 
Prof. A. E. Blount, while at the Colorado experiment station, made 
many wheat hybrids and obtained several improved varieties. Blount's 
Hybrid No. 15, a cross of Lost Nation with Sonora, has become a well- 
known race, giving excellent results in some States. Probably the 
most valuable work in wheat hybridization in this country has been 
done by A. N. Jones, of New York. Mr. Jones writes: "Most of 
my crossbreeds are from Russian and American varieties, with some 
blood from Mediterranean Longberry or offspring from these combi- 
nations." Of the sixteen or more hybrid wheat races introduced by 
Jones, several have become standard sorts. Winter Fife, which is 
extensively grown in Indiana, Ohio, and other places, is probably his 
best-known race. His Early Red Clawson, Early Genesee Giant, etc., 
are among our widely grown races. 

From 1888 to the present time Prof. William Saunders, director of 
the experimental farm, Ottawa, Canada, has been hybridizing wheats 
particularly to secure early ripening races. To accomplish this lie 



488 YEARBOOK OP THE DEPARTMENT OF AGRICULTURE. 

has sought to secure earliness and hardiness in the best American 
races by hybridizing them with various Russian sorts. Preston and 
Stanley, derived from Ladoga, a Russian sort, crossed with Red Fife, 
and Alpha, Percy, and Advance, derived from Ladoga crossed with 
White Fife, are proving valuable additions. Tests of Preston and 
Advance at the Minnesota experiment station have given good results. 
Professor Hays, of that station, says: "Preston is the most inter- 
esting and promising variety of wheat procured outside of the State, 
and it bids fair to be a strong rival of our best Fife and Blue Stem 
wheats." Besides his important work on selection mentioned above, 
Professor Hays has been in recent years conducting experiments in 
hybridization and has obtained results of the greatest promise. It is 
noteworthy that in this country the wheat hybrids thus far produced 
which have given valuable results are racial hybrids, in many cases 
very complex, including several different races. 

Oats. — No oat hybrids produced in this country have as yet become 
very important so far as the writers are informed, although some are 
of exceptional interest, as, for instance, Priiigle's Excelsior, a so-called 
hull-less oat produced by crossing the common Chinese Hull-less (Avena 
nuda) with the Excelsior, a race of the common oat. This remark- 
able hybrid is said to possess the strength and robust character of the 
common oat and to retain the peculiarity of the naked seed derived 
from the Chinese Hull-less. It was introduced about the year 1881, 
but does not appear to have proved satisfactory for general culture. 
Recently Garton Bros., of England, have introduced a similar " naked 
oat," which gives great promise of proving a valuable sort, particu- 
larly for the preparation of oatmeal and similar foods. Apparently 
hybridization in this line promises important results. 

IMPROVEMENT IN FLOWERS AND ORNAMENTAL PLANTS. 

In no plants has scientific plant breeding been carried further than 
in those grown for their flowers or for ornamental purposes. Growers 
of such plants are compelled to produce new and striking varieties 
and races, and so must take advantage of all available methods. 

An interesting example of the result of continuous selection is the 
Blanche Ferry sweet pea, which resulted from over twenty-five years 
of selection from the old Painted Lady, in northern New York. In 
successive years the plants gradually became more stocky and com- 
pact, until after ten or twelve years they needed no outside support. 
From the Blanche Ferry there have arisen independently at least two 
of the dwarf varieties known as "Cupids." These arose as seedling 
sports and soon became very widely diffused. 

Probably in no other plants has hybridization given such marked 
results as in those cultivated for their flowers. This is due largely to 
the fact that in such plants variation of form and color of flowers are 
the greatest desiderata, and such modifications are most easily obtained 

L«fC. 



PROGRESS OF PLANT BREEDING. 489 

by hybridizing different-colored species, varieties, etc. Orchids, 
roses, begonias, chrysanthemums, cannas, and many other of our 
common flowers have been crossed, and recrossed until it is f requently 
impossible to determine their origin. In this country probably the 
most attention has been given to roses, carnations, and chrysanthe- 
mums. It is to hybridization, directly or indirectly, that Ave are 
indebted for almost all the beautiful forms of these flowers. By the 
introduction of foreign species and their utilization in hybridization 
with those already in cultivation, new and almost totally different 
strains are frequentty produced. As an illustration of this may be 
mentioned the important results that have been produced by the 
recent introduction of the hardy roses Rosaritgosa and R. ivichuraiana 
and their hybridization with our common varieties of roses. Manda 
says : " By crossing Rosa ivichuraiana with greenhouse teas the result 
is astonishing, as the plants are not only hardy, but retain their foli- 
age during the winter. Thus a new race of evergreen roses has been 
added to our collection, and promises to be the beginning of a new 
and useful class." 

Advantage has been taken of still another principle in growing- 
plants of this class. It sometimes occurs that certain buds give rise 
to branches that vary abnormally and produce flowers or leaves of a 
different color or shape from those borne on the rest of the plant. 
These so-called bud sports can often be perpetuated, and thus give 
rise to new varieties. In this way many of the cut-leaved forms of 
various ornamental plants have originated. Perhaps the most strik- 
ing examples of the production of new sorts by bud sporting are found 
in certain plants, such as the chrysanthemum, rose, carnation, etc. 
Many of these are sports merely in color, but in some cases even the 
form of the flowers and of the leaves is different. It is said that within 
the last ten years there have been over fifty cases of new varieties of 
chrysanthemums originated as bud sports. 

IMPROVEMENT IN NUTS. 

Among the native nuts, probably the chestnut and the pecan are the 
only ones that have received much attention from plant breeders, 
though Burbank has given some care to improving the walnut. A few 
varieties of chestnuts have been obtained by the selection of wild trees 
of desirable quality, though but little more than this has been done. 
The pecan, however, has received more attention. In its wild state, it 
varies very greatly in its characteristics, and this has led to the selection 
of a number of varieties from wild trees because of some special quality, 
as thinness of shell, small amount of corky substance between the 
halves of the kernel, productiveness, size, or other good qualities. 
One of the best known of such selected pecans is the Frotscher, the 
original tree of which is still standing in Louisiana, and is probably 
over two hundred years old. Within the last fifty years many growers 



490 YEARBOOK OF THE DEPARTMENT OF AGRICULTURE. 

have been planting the nuts of this and other varieties and selecting 
from among the seedlings thus produced those with the best quali- 
ties. PI. XXXVIII, fig. 2, shows the variations and the possibilities 
of improvement when careful selection is exercised. The systematic 
improvement of the pecan, however, has just begun. 

IMPROVEMENT IN COTTON. 

The history of sea-island cotton is extremely interesting, as it 
serves as an example of the possibility of adapting a tropical plant to 
the conditions of culture in temperate regions. About 1785 seeds of 
this cotton were brought to Georgia from the Bahamas. Notwith- 
standing the good care they received and the mild winter, the plants 
were killed down, but they came up again from the roots, and with this 
start succeeded in ripening a few seeds before the first frost in the fall. 
The earliest of these seeds were sown in turn, and by continuing this 
process of selection the flowering period became earlier and earlier, 
until now the plants ripen a large proportion of their seeds before 
frost, even along the coasts of the Carolinas. Besides striving to 
obtain earlier maturing sorts, very careful selection has for years been 
made Avith a view of increasing the length, fineness, and strength of 
the staple. This selection is regularly practiced by all intelligent 
growers and to-day it may be regarded as one of the necessary cultural 
methods. Every year a special patch of cotton is grown from selected 
seed; the plants in this patch are examined very carefully and the 
seed of the best individuals retained for planting a similar patch 
the next year, the seed of the remaining plants being used to plant 
the general crop. Under such continuous and vigorous selection the 
length and fineness of the fiber have gradually increased, until it is now 
recognized as superior to that grown anywhere else in the world and 
commands the highest price in the market. 



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